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NOTE Eur. J. Entomol. 104: 139–143, 2007 http://www.eje.cz/scripts/viewabstract.php?abstract=1207 ISSN 1210-5759

Increased migration of linked to climate change

TIM H. SPARKS1, ROGER L.H. DENNIS1,2, PHILIP J. CROXTON1 and MARTIN CADE3

1NERC Centre for Ecology and Hydrology, Monks Wood, Abbots Ripton, Huntingdon, Cambridgeshire PE28 2LS, UK; e-mail: [email protected] 2Institute for Environment, Sustainability and Regeneration, Mellor Building, Staffordshire University, College Road, Stoke on Trent, ST4 2DE, UK 3Portland Bird Observatory, The Old Lower Light, Portland Bill, Dorset DT5 2JT, UK

Key words. Lepidoptera, , , migration, temperature response, climate change

Abstract. The number of of migratory Lepidoptera (moths and butterflies) reported each year at a site in the south of the UK has been rising steadily. This number is very strongly linked to rising temperatures in SW . It is anticipated that further cli- mate warming within Europe will increase the numbers of migratory Lepidoptera reaching the UK and the consequences of this invasion need urgent attention.

INTRODUCTION treated in the sense of Williams (1965). Since records were made in a semi-quantitative way (for example, a text description Evidence is steadily accumulating on the impact of climate of numbers or duration), we have reduced information to binary change on natural systems (Walther et al., 2002; Root et al., records of presence and absence in each year for each species. 2003). Documented changes in Lepidoptera, usually butterflies, Preliminary analysis examining temperatures in the UK and as a consequence of rising temperatures are focussed on the continental Europe suggested greatest correlations with tem- northern temperate zone and include advanced phenology (Roy peratures in SW Europe. Mean monthly temperature anomalies & Sparks, 2000; Forister & Shapiro, 2003; Stefanescu et al., (differences from the 1961–1990 average) were obtained from 2003, Dell et al., 2005), changes in morphology, widening the 5° gridded CRUTEM2v dataset (www.cru.uea.ac.uk). Tem- habitat base, increased population size, and altitudinal (Kon- peratures for SW Europe were approximated by the average of viþka et al., 2004; Davies et al., 2005) and geographical range the four grid boxes 35–45°N 10–0°W in and southern shifts (Parmesan et al., 1999). These may partly compensate for France. population declines arising from habitat degradation and poten- Least squares regression was used to compare the total tial extinctions (Warren et al., 2001; Thomas et al., 2004; Fox et number of migratory species recorded each year with tempera- al., 2006). Little has been published to date on the consequences ture anomalies. For the analysis of the presence/absence of indi- of climate change on the migration of , other than pest vidual species using logistic regression it is only possible to species (Cannon, 1998; Robinson et al., 2005). Work on historic examine species with both present and absent states. Hence, data (up to 1962) suggested increased migration of Lepidoptera only species recorded as present in between five and 19 years into Britain in years when temperatures in mainland Europe were compared to monthly temperature anomalies between were higher (Sparks et al., 2005) but little research has been January and September using forwards selection binary logistic done on contemporary data. regression. In this short paper we examine incidence of, and the influence of temperature on, the migration of Lepidoptera into a site on RESULTS the south coast of the UK. The migration routes and origins of A grand total of 75 species of migratory Lepidoptera were these species are largely unknown but, broadly speaking, at this recorded at Portland between 1982 and 2005 (Table 1). location the migrants will inevitably originate from south- The annual numbers of species of migratory Lepidoptera western Europe. varied between eight and 43 (mean 25) and increased signifi- 2 MATERIAL AND METHODS cantly by an average of 1.34 ± 0.15 species/annum (Fig. 1, R = 79.3%, F1,22 = 84.28, p < 0.001). Numbers of migratory species Between 1982 and 2005, records of the incidence of Lepidop- were positively related to temperature anomalies averaged over tera from both light trapping and observations during the day March to July and suggested a 1°C increase in temperature was have been taken in the cliff-top garden (ca. 0.2 ha) at the Port- associated with an additional 14.4 ± 2.4 migrant species (Fig. 2, 2 land Bird Observatory, Dorset, UK (50.55°N, 2.44°W) situated R = 61.9%, F1,22 = 35.79, p < 0.001). The March to July mean at the southern end of a 9 km headland extending into the Eng- temperature anomaly was also rising within this period (regres- 2 lish Channel. Recorder effort was approximately constant sion b = 0.058 ± 0.012, R = 50.2%, F1,22 = 22.22, p < 0.001). throughout the period. Recording takes place throughout the The predominantly positive nature of temperature anomalies year except on days of exceptional wind or very low winter tem- shown in Fig. 2 emphasises that current temperatures are peratures which experience has shown to be unproductive. warmer than the 1961–1990 average. Migratory status (full or partial) was determined according to There were 28 species with at least five contrasting binomial Emmet & Heath (1991) and thereby includes species that have states and their stepwise binary logistic regressions are summa- undergone overseas flight to British shores; the term is therefore rised in Table 2. Significant models were achieved for 22 of the

139 TABLE 1. The 75 species with some migratory status as indicated by Emmet & Heath (1991) and the number of years for which each were recorded at Portland between 1982 and 2005. Species with between five and 19 years of data were analysed individually using logistic regression. English name (if any) Scientific name Number of years recorded Hawk Agrius convolvuli 23 Dark Sword-grass Agrotis ipsilon 23 Silver Y Autographa gamma 22 Hummingbird Hawk Moth Macroglossum stellatarum 22 White-speck unipuncta 21 The Delicate Mythimna vitellina 21 Vestal sacraria 21 Red Admiral Vanessa atalanta 21 Painted Lady Cynthia cardui 20 Pearly Underwing Peridroma saucia 20 Clouded Yellow croceus 19 White Point Mythimna albipuncta 19 Small Mottled Spodoptera exigua 18 Scarce Bordered Straw armigera 17 Bordered Straw Heliothis peltigera 17 Rush Veneer noctuella 17 Gem obstipata 16 Diamond-Back Plutella xylostella 16 Rusty-dot Pearl ferrugalis 16 Jersey Tiger quadripunctaria 15 Cosmopolitan Mythimna loreyi 15 European Corn Borer Ostrinia nubilalis 13 Marbled-yellow Straw Pearl extimalis 12 11 Ni Trichloplusia ni 10 Great Dart Agrotis crassa 9 Scarce -tree Pearl unionalis 9 Knot-horn abietella 8 Striped Hawk Moth lineata 8 Isle of Wight Knot-horn oblitella 7 Rusty amplana 6 Monarch Danaus plexippus 6 Conobathra tumidana 5 Purple Marbled ostrina 5 Necklace Grass-veneer ocellea 5 Barred Red fasciaria 5 Palpita vitrealis 5 Alpine Grass-veneer alpinella 5 Hoary Footman caniola 4 Small Marbled 4 Bedstraw Hawk Hyles gallii 4 Straw Dot sericealis 4 Flame Brocade flammea 4 Blair’s Mocha puppillaria 3 Death’s Head Hawk Moth Acherontia atropos 2 Pale Shoulder lucida 2 Antigastra catalaunalis 2 Porter’s Rustic hospes 2 Red-headed erythrocephala 2 Many-lined Costaconvexa polygrammata 2 ramburialis 2 The Passenger algira 2 Pygmy Footman Eilema pygmaeola 2 bipunctella 2 140 TABLE 1 (continued). The 75 species with some migratory status as indicated by Emmet & Heath (1991) and the number of years for which each were recorded at Portland between 1982 and 2005. Species with between five and 19 years of data were analysed individually using logistic regression. English name (if any) Scientific name Number of years recorded Old World Webworm Hellula undalis 2 Bloxworth Snout Hyphena obsitalis 2 Four-spotted Footman quadra 2 Tawny Wave rubiginata 2 Small Thistle Moth Tebenna micalis 2 Rosy Underwing electa 1 Golden Twin-spot Chrysodeixis chalcites 1 Crombrugghia laetus 1 The Nutmeg Discestra trifolii 1 Angle-striped Sallow paleacea 1 Long Tailed Blue boeticus 1 Gypsy Moth Lymantria dispar 1 Dewick's Plusia confusa 1 Radford's Flame Shoulder Ochropacha leucogaster 1 Clancy’s Rustic Platyperigea kadenii 1 Powdered Pearl pulveralis 1 Dark Mottled Willow Spodoptera cilium 1 Cypress Carpet Thera cupressata 1 Orache Moth Trachea atriplicis 1 Yellow-underwinged Pearl Uresphita polygonalis 1 Crimson Speckled Utetheisa pulchella 1

28 species. Of the 39 terms included in all models only four had form (altitude, distance, direction) that migration takes is negative coefficients, indicating that higher temperatures in SW closely related to atmospheric conditions (Wood et al., 2006). Europe were generally associated with greater incidence of The relationships we calculate above suggest an increase of 14 migratory Lepidoptera in the southern UK. species for each 1°C increase in temperatures in SW Europe. Thus, a relatively modest degree of warming could make a sub- DISCUSSION stantial difference to the number of migratory Lepidoptera The majority of migratory Lepidoptera to the south coast of reaching new territories, furthermore numbers could well be England originate from the south and will have to fly over a higher on continental land masses where no physical barriers minimum of 150 km of open sea to reach the recording site. exist (e.g. Konviþka et al., 2004). Migration of insects into Britain, as between all land areas, is a The UK has a sparse fauna relative to its land area, persistent feature. It is considered to be the outcome of repro- thought to be a consequence of its island status and geological ductive bet-hedging – spreading breeding effort in space and history (Dennis, 1993). Even so, some 89 moth species alone time over a range of environmental conditions (Loxdale & Lushai, 1999; Holland et al., 2006, but see Dennis, 1993); the

Fig. 2. The relationship between the number of migrant Lepi- doptera species recorded each year at the Portland Bird Obser- Fig. 1. The number of migrant Lepidoptera species recorded vatory, UK and mean March–July temperature anomalies in SW each year at the Portland Bird Observatory, UK. Europe, see text for details.

141 TABLE 2. Stepwise logistic regression on nine monthly mean resident species, on their resources, but can also include species temperatures (January–September) for the 28 species which had of conservation concern over wider regions (e.g. , at least 5 years in an opposite state (present or absent). Months Pittaway, 1993). The threat imposed by migratory species are indicated by numerals (1 = January etc.) and are displayed relates not just to their mobility but to their adaptability which is in the order of entry. All coefficients were positive except believed to be closely linked to their mobility; migratory species where indicated by “(–)”. The deviance (G) and the overall may be among the most adaptable of species (Cannon, 1998). model significance (P) are also shown. For this reason, they may represent a competitive threat to resi- dent species which typically have lower mobility and are more Months included Species n G P specialised in habitat requirements. in model The possible consequences of a climate induced change in Colias croceus 19 7, 4 15.70 <0.001 burgeoning migrant insects, with potentially serious conse- Mythimna albipuncta 19 5 10.41 0.001 quences for health, agriculture and conservation of resident taxa, Spodoptera exigua 18 None requires immediate attention. Heliothis armigera 17 5, 1 11.40 0.003 REFERENCES Heliothis peltigera 17 5 5.80 0.016 CANNON R.J.C. 1998: The implications of predicted climate Nomophila noctuella 17 7, 2 23.84 <0.001 change for insect pests in the UK, with emphasis on non- 16 5 4.28 0.039 indigenous species. Global Change Biol. 4: 785–796. Plutella xylostella 16 8, 9 (–) 11.42 0.003 CHIN T. & WELSBY P.D. 2004: Malaria in the UK: past, present, and future. Postgrad. Med. J. 80: 663–666. 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